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ANKYLOSAURIDAE (DINOSAURIA) from MONGOLIA -.: Palaeontologia Polonica

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TERESA MARYANSKA ANKYLOSAURIDAE (DINOSAURIA) FROM MONGOLIA (plates 19-36) Abstract. - The paper deals with Asian Ank ylosauria, assigned to the family Ankylosauridae, BROWN. New material collected by the Polish-Mongolian exped itions to Mongolia in three Late Cretaceous formations (Djadokhta, Barun Goyot and Nernegt) is described. Two new genera and species: Saichania chulsanensis and Tarchia kielanae, both from , the ?Campanian Barun Goyot Formation, are described. Revised diagnoses of all Mongolian ankylosaurid species are given. An ossified conchal system in nasal cavities of Saichania chulsanensis, a unique feature among the extinct lower tetrapods is described. The postfrontal, epipterygoid, and additional ossifications in the antorbital wall closing the orbit in the front, as well as four pairs of secondary ossifications of dermal origine, which cover partly the skull bones, are found in the skull of Pinacosaurus grangeri and Saichania chulsanensis. Presented reconstruction of the hyoid apparatus of Saichania chulsanensis differs from that known in Psittacosaurus and Protoceratops and shows similarities to that of Squamata. In the pectoral girdle of Saichania chulsanensis the fusion of the coracoid with the transverse process of the first dorsal vertebra is stated. A morphological (and possibly also phylogenetical) sequence of Mongolian Ankylosauridae, beginning with an unnamed Early Cretaceous specimen from Khovboor, through Talarurus, Pinacosaurus, Saichania to " Dyoplosaurus" is presented. The position of Ankylosauria within the Ornithischia is discussed. It is presumed that the quadrupedality in Ankylosauria is primary. The origin of Ankylosauria from Pseudosuchia, independent from the line of bipedal and cursorial Ornithopoda seems possible. Common origin of Ankylosauria and Stegosauria is suggested. Contents Page Introduction . 86 Material . .. 93 Measurements. 95 General remarks on the systematics 97 Check list of Asian Ankylosauria 98 Descriptions . 99 Suborder Ankylosauria . 99 Family Ankylosauridae, BROWN 99 Genus Talarurus MALEEV . 99 Talarurus plicatospineus MALEEV . 100 Talarurus disparoserratus (MALEEV) 100 Genus Pinacosaurus GILMORE 101 Pinacosaurus grangeri GILMORE 101 Genus Saichania nov. 103 Saichania chulsanensis sp.n. 103 Genus Tarchia nov. 105 Tarchia kielanae sp.n, 108 Anatomy . 108 Osteology of the skull 109 Individual bones . 109 Accessory elements. 118 Bony nasal structures 119 Remarks on the nares and nasal cavities 122 86 TER ESA MARYANSKA Osteology of the postcranial skeleton. 124 Vertebral column and ribs . 124 Pectoral girdle and fore limb . 127 Pelvic girdle and hind limb .. 130 Dermal armour .. ... 131 Geographic and stIatigraphic distribution ot Ankylosauria in Asia 132 Evolution of Asiatic Ankylosauridae . 134 Position of Ankylosauria within Ornithischia . 136 Habits of Ankylosauridae 139 References .. 140 INTRODUCTION Ankylosauria from the Cretaceous of Asia have been studied only preliminarily. A mono­ graph of COOMBS (MS) devoted to ankylosaurs contains few data about Asiatic representatives. The first note on the occurrence of the armored dinosaurs in Asia come from MATLEY (1923) who has described Lametasaurus indicus from the Cretaceous of Jubbulpore (Lameta Beds). Assignment of Lametasaurus to the Ankylosauria has been questioned (CHAKRAVARTI, 1935). Notes by WIMAN (1929) and GILMORE (1933a) pertain to the occurrence of remains of the true Ankylosauria in China. These fragmentary findings did not allow more precise identification. Pinacosaurus grangeri GILMORE described on the basis of a deformed skull (AMNH 6523) from the Djadokhta Formation of Shabarakh Usu (Bayn Dzak) was the first Asiatic species undoubtedly belonging to Ankylosauria (GILMORE, 1933b). In 1935 YOUNG described Pina­ cosaurus ninghsiensis from the Cretaceous of.Ninghsia (China). In the USSR RIABININ (1939) has found ankylosaufs in the Upper Cretaceous near Tashkent. It seems also probable that the remains descr.bed by him as possible Ceratopsia (I.e., table 11, figs I, la, 4, 4a; table 12, figs 4,5; table 14, figs 2, 2a) belong also to the Ankylosauria. On the basis of incomplete and badly preserved materials found during 1930-1931 by the Sino­ Swedish expeditions, BOHLIN (1953) reported the occurrence of the ankylosaurs in several sites of Cretaceous age in the eastern Kansu and has described three new genera and species: Sauroplites scutiger, Heishansaurus pachycephalus and Peishansaurus philemys, as well as a new genus and species Stegosaurides excavatus assigned to ?Stegosauria. MALEEV 1 (1956), ROMER (1966), STEEL (1969) and others assigned Stegosaurides excavatus to the Ankylosauria. Not all species established by BOHLIN can be regarded as valid because of their incomplete type specimens. Data pertaining to occurrence of Ankylosauria in China are supplemented by the reports of the 1960 Sino-Soviet Palaeontological Expeditions (ROZHDESTVENSKY, 1961a, 1961b). Two skeletons of ankylosaurs have been excavated near Maortu (Alashan). This material so far undescribed is housed in Peking. Occurrence of ankylosaurs has also been noted in Inner Mongolia (Young, 1964) and in Nanhsiung (Kwantung) in the Nanhsiung Formation of Late Cretaceous age (YOUNG, 1965; CHENG et al., 1973). Abundant and well preserved remains of ankylosaurs have been excavated from the Upper Cretaceous of Mongolia by Soviet expeditons during 1946-1949 (MALEEV, 1952a, 1952b, 1952c, 1954b, 1956). Remains from Bayn Shireh (Bayn Shireh svita, see p. 132) described by MALEEV (1952b) as Talarurus plicatospineus include numerous specimens housed in the Pa­ laeontological Institute in Moscow. MALEEV (1952a) established a new family, the Syrmosauridae (regarded herein as a synonym of the Ankylosauridae) with one genus Syrmosaurus (type species Syrmosaurus viminicaudus) from the Djadokhta Formation. MALEEV (1952b) has also described Syrmosaurus disparoserratus from Sheeregeen Gashoon (Bayn Shireh svita) and Syrmosaurus sp. from Oshi. • Referred to in previous publications as MALEYEV. ANK.YLOSAURIDAE FROM MONGOLIA 87 The new ankylosaurs were discovered in the Upper Cretaceous of Mongolia by the Polish­ Mongolian Palaeontological Expeditions of 1963-1971 (KIELAN-JAWOROWSKA & DOVCHIN, 1969; KIELAN-JAWOROWSKA& BARSBOLD, 1972; MARYANSKA 1970, 1971). A rich and interesting collection of ankylosaurs was discovered by the Soviet-Mongolian Expeditions to Mongolia in 1970-1971. The latter collection contains Upper Cretaceous forms form e.g. Alag Teg (TVERDOCHLEBOV & TSYBIN, 1974) and Khermeen Tsav (KRAMARENKO, 1974) as well as Lower Cretaceous forms from the vicinity of Khovboor (KRAMARENKO, l.c.) and (?) Khuren Dukh (KALANDADZE & KURZANOV, 1974). Fragmentary undescribed remains of ankylosaurs from several sites in Kazakhstan (e.g. Shakh-shakh) housed in the Laboratory of Palaeobiology of Kazakh Academy of Sciences in Alma Ata and remains from Yakutia (USSR) housed in the Palaeontological Museum in Moscow complete the collection of ankylosaurs from Asia (see table 1 and text-fig. 1). The present paper is devoted to ankylosaurs collected in the Mongolian People's Republic by Polish-Mongolian Palaeontological Expeditions in 1963-1971, and housed in the Institute of Paleobiology of the Polish Academy of Sciences in Warsaw and in Geological Institute Section - i :90~ ---- \700 '- 60' /'0' I ! I 027 22 Fig. 1 Diagramatic map of Asia showing the localities in which ankylosaurs were found. 1 - Bayn Dzak, 2 - Alag Teg, 3 ­ AItan Via, 4 - Nemegt, 5 - Khulsan, 6 - Khermeen Tsav, 7 - Sheeregeen Gashoon, 8 - Khovboor, 9 - Bayn Shireh, 10 - Baga Tarjach, 11 - Baysheen Tsav, 12 - NW Ninghsia, 13 - Chia-yii-kuan, 14 - Hui-hui-pu, 15­ Ehr-hia wu-kuan, 16 - Maortu, 17 - Tsondolein Khuduk, 18 - Tebch, 19 - Iren Dabasu, 20 - Khuren Dukh, 21 - Oshi Nur, 22 - Jubbulpore, 23 - Alym Tau, 24 - Shakh-shakh, 25 - Djara Khuduk, 26 - Laiyang, 27- Nanhsiung, 28 - Kempenday. 8R TERESA MARYANSKA Ta Ankylosaurian Ide n t i f i c a t i o n Locality (forexpl an ati on Mu seum cat. no of numbers see origin al I in this paper Text-fig. 1) " Dyoplosaurus" gigantells PIN, undescribed speci- Khermen Tsav (6) MALEEY men , no catalogue number Dy oplosaurus cf. giganteus (MA- ZPAL MgD-I/43, housed RYANSKA, 1970) "Dyoplosaurus" giganteus no w in GISPS, VIan Bator Allan Via (3) M ALE EY Dyoplosaurus sp. (MARYANS KA, ZPAL MgD-I/42, a nd 1970) MgD-I/49, MgD-I/11 3 Dyoplosaurus giganteus MALE EY " Dyoplosaurus" giganteus PIN 551-29 Nemegt (4) (MALEEY, 1956) MALE EY . Pinacosauru s (KRAMARENKO, PIN, undescribed speci- 1974) men , no ca ta log ue number Saichania chulsan ensis gen. Khermeen Tsav II (6) n., sp. n. ankylosaurid (KI ELAN-JAwOROW- ZPAL MgD-I/114 SKA & BARSBOLD, 1972) ankylosaurid (KIELAN-JAwOROW- Sa ichania chulsanensis gen . GI SPS 100 /151 hol ot ype SKA & BARSBOLD, 1972) n., sp. n. Khulsan (5) ankylosaurid (KI ELAN'-JAWOROW- Tar chia kielanae gen. n., ZPAL MgD-I/111 hol otype SKA & BARSBOLD, 1972) sp. n. - Pinacosaurus ninghsiensis In st. Vert. Paleont., Pek ing , Pinacosaurus grangeri NW of Ninghsia (12) YOUNG (YOUNG, 1935) GILMORE no cat alogue number PIN, undescribed speci- ankylosaurs KUROCHKIN & K A- Pinaco saurus grangeri men s, no ca ta log ue num- Alag Teg (2) LANDADZE 1970) GILMORE bers Pinacosaurus grangeri GILMORE AMNH 6523 holot ype (GILMORE , 1933) Syrmosaurus viminicaudus M A- PIN 614 LEEY (M ALEEY, 1952) Bayn D zak (1) Pinacosaurus grangeri GI L- ZPAL M gD -I1/7 MORE ZP AL MgD-I1 /9 Pinacosaurus grangeri GI LMORE ZPAL MgD-II/31 (MARYANSKA, 1971) ZP AL MgD-II/32 ZPAL MgD-II/27
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    Dinosaur Species List E to M E F G • Echinodon becklesii • Fabrosaurus australis • Gallimimus bullatus • Edmarka rex • Frenguellisaurus • Garudimimus brevipes • Edmontonia longiceps ischigualastensis • Gasosaurus constructus • Edmontonia rugosidens • Fulengia youngi • Gasparinisaura • Edmontosaurus annectens • Fulgurotherium australe cincosaltensis • Edmontosaurus regalis • Genusaurus sisteronis • Edmontosaurus • Genyodectes serus saskatchewanensis • Geranosaurus atavus • Einiosaurus procurvicornis • Gigantosaurus africanus • Elaphrosaurus bambergi • Giganotosaurus carolinii • Elaphrosaurus gautieri • Gigantosaurus dixeyi • Elaphrosaurus iguidiensis • Gigantosaurus megalonyx • Elmisaurus elegans • Gigantosaurus robustus • Elmisaurus rarus • Gigantoscelus • Elopteryx nopcsai molengraaffi • Elosaurus parvus • Gilmoreosaurus • Emausaurus ernsti mongoliensis • Embasaurus minax • Giraffotitan altithorax • Enigmosaurus • Gongbusaurus shiyii mongoliensis • Gongbusaurus • Eoceratops canadensis wucaiwanensis • Eoraptor lunensis • Gorgosaurus lancensis • Epachthosaurus sciuttoi • Gorgosaurus lancinator • Epanterias amplexus • Gorgosaurus libratus • Erectopus sauvagei • "Gorgosaurus" novojilovi • Erectopus superbus • Gorgosaurus sternbergi • Erlikosaurus andrewsi • Goyocephale lattimorei • Eucamerotus foxi • Gravitholus albertae • Eucercosaurus • Gresslyosaurus ingens tanyspondylus • Gresslyosaurus robustus • Eucnemesaurus fortis • Gresslyosaurus torgeri • Euhelopus zdanskyi • Gryponyx africanus • Euoplocephalus tutus • Gryponyx taylori • Euronychodon
  • Reidentification of Avian Embryonic Remains from the Cretaceous of Mongolia

    Reidentification of Avian Embryonic Remains from the Cretaceous of Mongolia

    RESEARCH ARTICLE Reidentification of Avian Embryonic Remains from the Cretaceous of Mongolia David J. Varricchio1*, Amy M. Balanoff2, Mark A. Norell3 1 Department of Earth Sciences, Montana State University, Bozeman, Montana, 59717, United States of America, 2 Department of Anatomical Sciences, Stony Brook University School of Medicine, Stony Brook, NY, 11794, United States of America, 3 Division of Paleontology, American Museum of Natural History, New York, NY, 10024, United States of America * [email protected] Abstract Embryonic remains within a small (4.75 by 2.23 cm) egg from the Late Cretaceous, Mongo- lia are here re-described. High-resolution X-ray computed tomography (HRCT) was used to digitally prepare and describe the enclosed embryonic bones. The egg, IGM (Mongolian In- stitute for Geology, Ulaanbaatar) 100/2010, with a three-part shell microstructure, was originally assigned to Neoceratopsia implying extensive homoplasy among eggshell char- acters across Dinosauria. Re-examination finds the forelimb significantly longer than the hindlimbs, proportions suggesting an avian identification. Additional, postcranial apomor- phies (strut-like coracoid, cranially located humeral condyles, olecranon fossa, slender radi- OPEN ACCESS us relative to the ulna, trochanteric crest on the femur, and ulna longer than the humerus) identify the embryo as avian. Presence of a dorsal coracoid fossa and a craniocaudally Citation: Varricchio DJ, Balanoff AM, Norell MA (2015) Reidentification of Avian Embryonic Remains compressed distal humerus with a strongly angled distal margin support a diagnosis of IGM from the Cretaceous of Mongolia. PLoS ONE 10(6): 100/2010 as an enantiornithine. Re-identification eliminates the implied homoplasy of this e0128458. doi:10.1371/journal.pone.0128458 tri-laminate eggshell structure, and instead associates enantiornithine birds with eggshell Academic Editor: Peter Dodson, University of microstructure composed of a mammillary, squamatic, and external zones.
  • EUOPLOCEPHALUS ANKYLOSAURUS TSAGANTEGIA GASTONIA GARGOYLEOSAURUS - -- PANOPLOSAURUS 7,8939, (6, - Edmontonla 1L,26,34)

    EUOPLOCEPHALUS ANKYLOSAURUS TSAGANTEGIA GASTONIA GARGOYLEOSAURUS - -- PANOPLOSAURUS 7,8939, (6, - Edmontonla 1L,26,34)

    THE UNIVERSITY OF CALGARY Skull Morphology of the Ankylosauria by Matthew K. Vickaryous A THESIS SUBMITED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE DEPARTMENT OF BIOLOGICAL SCIENCES CALGARY, ALBERTA JANUARY, 2001 O Matthew K. Vickaryous 2001 National Library Bibliothéque nationale I*l of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395, rue Wellington Ottawa ON KIA ON4 Ottawa ON KIA ON4 Canada Canada Your Rie Votre rrlftimce Our fite Notre dfbrance The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/film, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Abstract The vertebrate head skeleton is a fundamental source of biological information for the study of both modern and extinct taxa. Detailed analysis of structural modifications in one taxon frequently identifies developmental and / or functional features widespread amongst a more inclusive clade of organisms.